Bonn zoological Bulletin Volume 57 Issue 2 pp. 231–240 Bonn, November 2010

Another new Cophixalus species (Amphibia: Anura: ) from western

Rainer Günther Museum für Naturkunde Berlin, Leibniz Institute for Research on Evolution and Biodiversity at the Humboldt-University, Invalidenstr. 43, D-10115 Berlin, Germany

Abstract. Based on external morphological, anatomical, bioacoustic, and molecular traits, a new species in the micro- hylid genus Cophixalus is described. The new species was discovered in the Fakfak Mountains, northwestern corner of the , Papua Province, . The new taxon is most closely related to the sympatric Cophixalus tetzlaffi. It differs from that species in several morphological traits, but primarily by its advertisement call: the new species utters a single peeping note with a mean duration of less than 200 milliseconds, whereas the advertisement call of C. tetz- laffi consists of three to four notes, with a mean note duration of more than 400 milliseconds. Molecular data (mitochon- drial 16S rRNA) support the determination of the specific distinctness of the new species. Key words. Anura, , Cophixalus, new species, New Guinea.

INTRODUCTION MATERIAL AND METHODS

Fifty-one species in the microhylid genus Cophixalus Most were collected at night after locating them by are known at present (Frost 2010). Of these, 14 occur in their advertisement calls. Some specimens were pho- north-eastern Australia, 30 in , three tographed in life the next day and all specimens were are known only from the Papua Province of Indonesia, anaesthetized with chlorobutanol and subsequently fixed three are recorded from both Papua New Guinea and the in 2 % formalin. Tissue probes from thigh muscle were Papua Province of Indonesia, and one species seems to be taken from some frogs and stored in about 96 % ethanol endemic to the Island of Halmahera, located about 300 km to enable DNA sequencing, before fixing the in west of the western tip of New Guinea. Although many formalin. All specimens were transferred to 75 % ethanol new Cophixalus species are expected to be described al- later in the Berlin museum. One specimen was cleared and so from , the distribution centre of stained as an osteological preparation according to a the genus seems to be clearly in eastern New Guinea and method modified from Dingerkus & Uhler (1977). north-eastern Australia. Cophixalus montanus has been known since 1895 from Halmahera, and the detection of The following measurements were taken with a digital cal- three new species in the western part of New Guinea (on liper (> 10 mm) or with a binocular dissecting microscope Yapen Island, on the Wandammen Peninsula, and on the fitted with an ocular micrometer (< 10 mm) to the near- Bomberai Peninsula) came as a surprise (Günther 2003, est 0.1 mm: 2006). SUL – snout-urostyle length: from tip of snout to distal tip of urostyle-bone; SUL is about one to two mm Here I describe another new species from the Fakfak shorter than the snout-vent length (SVL). As the Mountains on the Bomberai Peninsula (located on the measurement error is higher in the latter, I prefer to “throat” of the Vogelkop) found during an expedition in use the former. In general, both measurements are September 2008. Moreover, a population of Cophixalus more or less identical and are used interchangeably tridactylus and one specimen of a second undescribed in this paper. species was observed there. Consequently, at least four TL – tibia length: external distance between knee and an- Cophixalus species occur syntopically in the middle and kle; higher elevations (400–1000 m above sea level, a.s.l.) of TaL – length of tarsus: external distance, tarsal and ankle the Fakfak Mountains. joints held at a right angle;

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Fig. 1. Map of the western part of New Guinea with type lo- cality (1).

T4L – length of fourth toe: from tip of toe to proximal end of inner metatarsal tubercle; T4D – transverse diameter of disc of fourth toe; F3L – length of third toe; F3D – transverse diameter of disc of third finger; F1D – transverse diameter of disc of first finger; T1D – transverse diameter of disc of first toe; HL – head length: from tip of snout to posterior margin of tympanum; HW – head width, taken in the region of the tympana; SL – snout length: from an imaginary line that connects the centres of eyes to tip of snout; END – distance from anterior corner of orbital opening to centre of naris; IND – internarial distance between centres of nares; Fig. 2. Holotype of Cophixalus monosyllabus sp. n. head in lateral view (above); head in dorsal view (below). ED – eye diameter: from anterior to posterior corner of orbital opening; TyD – horizontal diameter of tympanum. Advertisement calls were recorded under natural condi- direct line NNE of Fakfak town, near the Fakfak-Kokas tions with a Sony Digital Audio Tape (DAT) Walkman road, Bomberai Peninsula (neck of Vogelkop), Papua TCD-D 100 and a Sennheiser microphone MKE 300 and Province, Indonesia, 2°53’S and 132°18’E, elevation 500 analysed with Avisoft-SAS Lab Pro software. All speci- m a.s.l., 9 September 2008 (Fig. 1). mens are currently stored in the Museum für Naturkunde Berlin (ZMB) and bear registration numbers of that insti- Paratypes. ZMB 74994 (FN: RG 7889), ZMB 74995 tution. Part of the type series will later be transferred to (FN: RG 7890), ZMB 74996 (FN: RG 7912), ZMB 74997 the Museum Zoologicum Bogoriense (MZB). (FN: RG 7915), ZMB 74998 (FN: RG 7916), ZMB 74999 (FN: RG 7926), ZMB 75000 (FN: RG 7927), ZMB 75001 (FN: RG 7951), ZMB 75002 (FN: RG 7952). ZMB 74997 RESULTS AND DISCUSSION is now an osteological preparation. All nine paratypes are males. They were collected from 9 to 14 September 2008 Cophixalus monosyllabus sp. n. along the Fakfak-Kokas road in the southern part of the Fakfak Mountains, at elevations of from 400 to 700 m Holotype. ZMB 74993 (field number, FN: RG 7888) ; a.s.l. Collectors were R. Günther, M. Kapisa, and A. and adult male collected by R. Günther and A. Piahar 6 km F. Piahar.

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Table 1. Body measurements and body ratios of the type series of Cophixalus monosyllabus sp. n. ZMB-No are the inventory numbers of the Museum für Naturkunde Berlin, FN are the field numbers of the author, SD indicates the standard deviation. ZMB 74993 is the holotype; ZMB 74997 is now an osteological preparation. All specimens are adult males. All measurements are in mm; abbreviations are explained in “Material and methods”.

ZMB-No 74993 74994 74995 74996 74997 74998 74999 75000 75001 75002 mean SD

FN 7888 7889 7890 7912 7915 7916 7926 7927 7951 7952 22.9 1.04

SUL 22.8 23.1 24.1 22.5 22.4 23.6 20.6 22.8 24.3 23.0 TL 10.9 11.7 11.4 11.5 11.3 11.5 10.1 11.7 12.3 11.7 TaL 7.4 7.2 7.1 6.8 6.9 7.2 7.2 7.6 7.3 7.3 T4L 11.5 11.6 11.7 11.6 11.2 11.1 10.5 11.1 12.5 10.9 T4D 1.3 1.25 1.4 1.0 1.3 1.3 1.0 1.2 1.35 1.25 T1D 0.7 0.65 0.75 0.5 0.7 0.6 0.45 0.7 0.6 0.6 F3L 6.0 6.4 6.8 6.4 6.1 6.5 5.5 5.9 6.8 6.0 F3D 1.4 1.4 1.7 1.45 1.5 1.3 1.25 1.5 1.5 1.4 F1D 0.6 0.45 0.50 0.50 0.5 0.4 0.45 0.5 0.5 0.5 HL 7.5 7.2 8.2 7.1 7.3 8.0 6.8 7.3 7.5 7.5 HW 8.5 9.0 9.1 9.0 9.5 9.8 8.5 9.0 9.6 9.6 SL 3.2 3.3 3.6 3.3 3.5 3.4 3.2 3.3 3.5 3.6 END 2.2 2.1 2.5 2.0 2.1 2.2 2.0 2.1 2.2 2.0 IND 2.3 2.5 2.5 2.3 2.2 2.4 2.2 2.5 2.5 2.25 ED 2.8 2.9 3.1 2.5 2.9 2.8 2.7 2.8 3.0 3.0 TyD 1.2 1.0 1.2 1.0 1.0 1.1 1.0 0.9 1.0 1.0

TL/SUL 0.48 0.51 0.47 0.51 0.50 0.49 0.49 0.51 0.51 0.51 0.50 0.051 TaL/SUL 0.32 0.31 0.29 0.30 0.31 0.31 0.35 0.33 0.30 0.32 0.31 0.017 T4L/SUL 0.50 0.50 0.49 0.52 0.50 0.47 0.51 0.49 0.51 0.47 0.50 0.016 F3L/SUL 0.26 0.28 0.28 0.28 0.27 0.28 0.27 0.26 0.28 0.26 0.27 0.009 F3D/SUL 0.061 0.060 0.071 0.064 0.067 0.055 0.061 0.066 0.062 0.061 0.063 0.004 F1D/SUL 0.026 0.019 0.021 0.022 0.031 0.017 0.022 0.022 0.021 0.022 0.021 0.002 T4D/SUL 0.057 0.054 0.058 0.044 0.058 0.055 0.049 0.053 0.056 0.054 0.054 0.004 T1D/SUL 0.031 0.028 0.031 0.022 0.031 0.025 0.022 0.031 0.025 0.026 0.027 0.004 HL/SUL 0.33 0.31 0.34 0.32 0.33 0.34 0.33 0.32 0.31 0.33 0.33 0.011 HW/SUL 0.37 0.39 0.38 0.40 0.42 0.42 0.41 0.39 0.40 0.42 0.40 0.018 HL/HW 0.88 0.80 0.90 0.79 0.77 0.82 0.80 0.81 0.78 0.78 0.81 0.043 END/IND 0.96 0.84 0.84 0.87 0.95 0.92 0.91 0.84 0.88 0.89 0.89 0.044 ED/SUL 0.123 0.125 0.129 0.111 0.129 0.119 0.131 0.123 0.120 0.130 0.124 0.006 TyD/ED 0.43 0.34 0.39 0.40 0.34 0.39 0.37 0.32 0.33 0.33 0.36 0.037 SL/SUL 0.140 0.143 0.149 0.147 0.156 0.144 0.155 0.145 0.141 0.156 0.148 0.006

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Diagnosis. With a snout-urostyle length of from 20.6 to 24.3 mm in ten adult males, the new species belongs to the middle-sized species of the genus. It is obviously a sis- ter species of the sympatric Cophixalus tetzlaffi and dif- fers from all other species in the same characters as the latter. The new species differs from C. tetzlaffi, among others, by its larger body size, its wider finger and toe discs, and its advertisement call which consists of only one peeping syllable (note) that lasts, on average, 196 millisec- onds (ms). In contrast, the advertisement call of C. tetzlaf- fi consists of three to four peeping notes, with a mean note length of more than 400 ms.

Description of the holotype. For measurements see Table 1. Head broader than long (HL/HW ratio 0.88), canthus rostralis roundish; loreal region straight; snout slightly pro- truding in profile (Fig. 2, above) and rounded in dorsal view (Fig. 2, below); horizontal eye diameter greater than eye-naris distance; borders of tympanum scarcely visible, its size less than half of the eye diameter (TyD/ED 0.43), no supratympanic fold; internarial distance slightly Fig. 3. Holotype of Cophixalus monosyllabus sp. n. ventral greater than distance between eye and naris (END/IND view of right hand (left); ventral view of right foot (right). 0.96); tongue large, posteriorly broadened and without posterior notch, its posterior and lateral margins free; a strongly serrated fold present in front of the pharynx; long

Fig. 4. Dorsolateral view of a more brownish coloured paratype of Cophixalus monosyllabus sp. n. (ZMB 74995).

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Fig. 5. Dorsolateral view of a more greyish coloured paratype of Cophixalus monosyllabus sp. n. (ZMB 74999).

slits on both sides of the tongue are entrances to a subgu- lar vocal sac. Legs moderately long, no webs between fin- gers or toes (Fig. 3); tips of fingers wider than tips of toes, first finger much smaller than other fingers, its tip only scarcely wider than the penultimate phalanx; relative length of fingers 3>2=4>1; third toe clearly longer than the fifth, tip of first toe slightly smaller than tip of the fifth toe, tips of remaining toes clearly wider than that of first and fifth toe; all finger and toe tips with terminal grooves; relative length of toes 4>3>5>2>1, all subarticular tuber- cles as well as metatarsal and metacarpal tubercles not or only scarcely developed. With exception of some tuber- cles on flanks, all dorsal, lateral, and ventral surfaces smooth.

Dorsum light brown and clearly demarcated against dark brown upper flanks, dorsal surfaces of legs non-uniform brown, chevron-shaped mark in scapular region, dorsal surface of snout lighter than remaining body; dorsal sur- faces of fingers and toes with yellowish, light brown, and dark brown pattern; lateral and dorsolateral flanks with longitudinal rows of blackish spots, a conspicuous black- ish spot present also above insertion of foreleg and behind eye; loreal region, tip of snout, and region below and be- Fig. 6. Ventral view of a paratype of Cophixalus monosylla- hind eye and underneath tympanum blackish (black face bus sp. n. (ZMB 74995). mask); ventral surface of forelegs yellowish with irregu-

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Fig. 7. Wave form (above) and spectrogram (below) of an advertisement call of Cophixalus monosyllabus sp. n..

lar dark brown flecks, ventral surface of hind legs also yel- lowish with brown flecks but the latter less intense than on forelegs (ventral skin and muscle tissue of the right thigh was removed for biochemical studies); belly, chest and throat yellowish with brown pigmentation, pigmen- tation most intense on throat and chest and least intense on abdomen; region around anal opening blackish and re- gion from behind eye, through tympanum, and up to up- per arm whitish.

Variation in the type series: Mensural variation for the type series is shown in Table 1. The basic colour and colour pattern elements of all preserved types are fairly uniform and very much resemble those of the holotype. Charac- teristic elements are a light brown dorsum, which is dif- ferentiated from dark brown upper flanks, a dark brown and irregularly pronounced interocular band, a dark chevron or W-shaped mark in the scapular region, a dark face mask, a blackish spot behind eye and above inser- tion of fore leg, a blackish throat which fades posterior- ly into a diffuse dark brown reticulum, and a pale dorsal surface of the snout which is the palest part of all the dor- sal surfaces. Only one specimen (ZMB 74996) has a whitish middorsal line from snout tip to anal opening and which continues on to the posterior thighs.

The basic colour in life varies from crème or grey to light brown. Dorsum rather uniform brownish (Fig. 4) or grey (Fig. 5); conspicuous is a blackish or dark brown chevron or W-shaped mark in the scapular region, an irregular dark brown interocular band between posterior parts of eyes, and a crème or light brown coloured dorsal part of snout. Fig. 8. Power spectrum of an advertisement call of Cophixa- Lower flanks are mostly lighter than the remaining later- lus monosyllabus sp. n. al areas (Fig. 4). Upper flanks may be of nearly the same

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Fig. 9. Habitat of Cophixalus monosyllabus sp. n. in the Fakfak Mountains on the Bomberai Peninsula, 700 m a.s.l. brown or grey colour as on the dorsum. Conspicuous are specimen in Fig. 5 disappeared completely in fixative. a blackish spot behind the eye, another blackish spot above Dorsal sides of legs similarly coloured as other dorsal and the foreleg insertion, and some blackish spots at the bor- dorsolateral body parts. Ventral sides of forearms crème- der between the dorsum and flanks. It is notable that the coloured, its anterior and posterior part covered with ir- dorsum in all preserved specimens is clearly lighter than regular dark spots. Throat and chest in all specimens dark- the upper flanks, whereas the dorsum and upper flanks in er than on the remaining ventral surfaces. These dark ven- most living specimens differed in colour only slightly. Lo- tral areas are solidly or discontinuously black or dark real region in all specimens entirely or predominantly brown. Abdomen and ventral sides of hind legs show black. In some specimens this black area continues to be- greater light areas covered by a more or less dense retic- low the eye and extends up to the upper arm, in others this ulum of grey-brown. Weakest pigmentation was common- black area ends below the eye. The inner margin of the ly on the posterior abdomen (Fig. 6). Iris yellow-red and “upper eyelid” is whitish in most specimens, this colour nerved by a dense net of blackish lines. merging in a broad and light postocular band. Osteology. One cartilage-bone preparation (ZMB 74997) While there are no or only a few tubercles in the preserved did not show remarkable differences from that of Cophix- specimens, most living specimens exhibited tubercles on alus tetzlaffi (see Günther 2003). the flanks and extremities (rarely on the dorsum). Many of these tubercles have a blackish base and a orange-red Vocalisation. Most calling activities were recorded dur- cap. Most, and the largest, tubercles are arranged in dor- ing rain and damp weather from dusk to 9 p.m. All calls solateral rows. The yellowish spot posterior of the were recorded at temperatures of approximately 21°C. chevron sign in ZMB 74999 (Fig. 5), which faded to a Calls are uttered in series lasting several minutes. The white spot in preservative, is obviously an exception. Or- shortest time between two successive calls was about 3 ange-red areas were also found on the forelegs of some s. Each call consists of a single unpulsed and finely tuned specimens. The fine whitish middorsal line in the living note (Fig. 7). Fifty-six calls (notes) from two males had

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Fig. 10. An undescribed Cophixalus species from the Fakfak Mountains, with a 16.9 mm snout-urostyle-length, which at first glance resembles Cophixalus misimae recently described by Richards & Oliver (2007) from Misima Island, Louisiade Archipela- go, Papua New Guinea. a mean length of 196 ms, with a minimum of 173 ms and only two m from one another. At favoured places about a maximum of 224 ms. Most notes start with a sharp in- ten males could be heard calling from one point. For bio- crease in amplitude, and the sound volume may remain geographical reasons it seems worthwhile to mention that constant during the entire note but may also change, with at elevations of between 500 and 700 m a.s.l., C. mono- the greatest sound volume mostly near the end. The end syllabus sp. n. occurs syntopically with C. tetzlaffi, C. tri- of the note occurs more gradually and its exact cessation dactylus, and another obviously new Cophixalus species is fairly difficult to identify (Fig. 7). The dominant fre- quency is approximately 2.8 kHz (Fig. 8), the fundamen- tal frequency is approximately 1.4 kHz, and the first (and most pronounced) upper harmonic band is at about 4.2 kHz.

Distribution. The new species lives on slopes and in val- leys of the southern part of the Fakfak Mountains. We found it along the Fakfak town-Kokas road at elevations of from 250 to 700 m a.s.l. Whether it also occurs in the northern part of the Fakfak Mountains remains to be de- termined.

Habitat and habits. Cophixalus monosyllabus sp. n. lives mostly in the understory (bushes, young trees, and herbs) of taller trees but also in shrubbery without a canopy cov- er (Fig. 9). The frogs perched mainly on or between liv- ing or dead leaves at heights of from one to three meters above the ground. The species is common: we heard sev- eral hundred males during a walk of three kilometres along Fig. 11. Bayesian inference phylogram of 16S rRNA. Numbers the Fakfak-Kokas road. Some males called at distances of on branches denote posterior probabilities.

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(Fig. 10). Ecological differences between these four species are the following: C. monosyllabus sp. n. occurs at from 250 to 700 m a.s.l. and its calling sites are at be- tween one and three metres above ground; C. tetzlaffi oc- curs at from 400 to 900 m a.s.l. (top of the mountains) and its calling sites are on structures up to one m above the ground; C. tridactylus was found at from 500 to 900 m a.s.l. and its calling sites are on the ground; and the ob- viously new species was found at 860 m a.s.l. in humus soil below the ground surface.

Etymology. The Latin word “monosyllabus” is derived from the Greek composite adjective “monosyllabos” meaning one syllable or monosyllabic, and refers to the advertisement call of the new species which consists of only one uniform note. I dedicate this new species to my dear colleague of many years, Prof. Dr. Wolfgang Böhme, to acknowledge his extraordinary contributions to herpetological science and on the occasion of his re- tirement from official service, although it is well known that Wolfgang is by no means monosyllabic but rather is very eloquent.

Molecular evidence. According to B. Stelbrink and T. von Rintelen (pers. comm., July 2010) DNA isolation and PCR were done using the protocol of Köhler & Günther (2008). Forward and reverse strands were aligned using Codon- Code Aligner v. 3.0.3 (CodonCode Corporation, Dedham, MA, USA) and corrected by eye. Sequences were aligned using MAFFT (Katoh & Toh 2008) and optimized using ALISCORE (Misof & Misof 2009). Phylogenetic analy- sis (Bayesian inference) was accomplished as conducted by Günther et al (2010).

The analysis of 480 base pairs of the 16S rRNA gene re- vealed that Cophixalus monosyllabus sp. n. is clearly a sister species of C. tetzlaffi and both are a sister clade of C. balbus (Fig. 11). C. tridactylus and C. humicola appear more distant in the molecular tree (see also Köhler & Gün- ther 2008), and indicate that the present genus Cophixalus most probably is polyphyletic. The genetic distance (un- corrected p-distance) between C. monosyllabus sp. n. and C. tetzlaffi is 4.3 % for the 16S rRNA gene.

Comparison with other species. Cophixalus monosylla- bus sp. n. is distinct from other Cophixalus species, des- cribed up to the year 2003, in the same characters as is C. tetzlaffi (Günther 2003). All 16 Cophixalus species des- cribed after 2003 (Hoskin 2004; Kraus & Allison 2006, 2009; Günther 2006; Richards & Oliver 2007) differ clearly from C. monosyllabus sp. n. in body size and al- so in their advertisement calls. The only species with Fig. 12. Box-Whisker-Plot of the ratio “diameter of disc of which C. monosyllabus sp. n. can be confused morpho- fourth toe/snout-urostyle-length” (F3D/SUL) in Cophixalus tetz- logically is C. tetzlaffi, especially as both species occur laffi (A) compared to that of Cophixalus monosyllabus sp. n. (B). syntopically.

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I compared the measurements of ten male C. monosyllabus REFERENCES sp. n. with that of eight male C. tetzlaffi and found the fol- Dingerkus G, Uhler LD (1977) Enzyme clearing of alcian blue lowing differences: with a mean body size (SUL) of 23.0 stained whole small vertebrates for demonstrating cartilage. mm (range 20.6–24.3 mm), C. monosyllabus sp. n. is Stain Technology 52: 229–232 somewhat larger than C. tetzlaffi (mean 21.4 mm, range Frost DR (2010) Species of the World: an online 19.5–22.6 mm), Student´s t-test revealed a significant dif- reference, version 5.4 (8 April 2010). American Museum of ference with t=2.98 and P=0.0046 in this character; C. Natural History, New York. Available from: monosyllabus sp. n. has significantly shorter tibiae than http://research.amnh.org/herpetology/amphibia/index.php Günther R (2003) First record of the microhylid frog genus C. tetzlaffi (mean of TL/SUL in C. monosyllabus sp. n. Cophixalus from western Papua, Indonesia, with descriptions 0.50, that in C. tetzlaffi 0.52, t= 3.39, P=0.0019); C. mono- of two new species. Herpetozoa 16 (1/2): 3–21 syllabus sp. n. has a longer third finger than C. tetzlaffi Günther R (2006) Two new tiny Cophixalus species with reduced (mean of F3L/SUL in the former 0.27, in the latter 0.26, thumbs from the west of New Guinea (Anura: Microhylidae). t=3.01, P=0.0041; C. monosyllabus sp. n. has a wider ter- Herpetozoa 19 (1/2): 59–75 minal disc on the fourth toe than C. tetzlaffi (mean of Günther R, Stelbrink B, Rintelen T von (2010). Oninia senglaubi, another new genus and species of frog (Amphibia, Anura, Mi- T4D/SUL in the former 0.054, in the latter 0.046, t=3.11, crohylidae) from New Guinea. Zoosystematics and Evolution P=0.0094); C. monosyllabus sp. n. has a wider terminal 86 (2): 245–256 disc on first toe than C. tetzlaffi (mean of T1D/SUL in the Hoskin CJ (2004) Australian microhylid frogs (Cophixalus and former 0.027, in the latter 0.018, t=5.63, P=0.00002); C. Austrochaperina): phylogeny, , calls, distributions monosyllabus sp. n. has a wider terminal disc on first fin- and breeding biology. Australian Journal of Zoology 52: 237– ger than C. tetzlaffi (mean of F1D/SUL in the former 269 Katoh K, Toh H (2008) Recent developments in the MAFFT 0.021, in the latter 0.015, t=5.36, P=0.00003) and, most multiple sequence alignment program. Briefings in Bioinfor- significantly, C. monosyllabus sp. n. has a wider terminal matics 9: 286–298 disc on the third finger than C. tetzlaffi (mean of F3D/SUL Köhler F, Günther R (2008) The radiation of microhylid frogs in the former 0.063, and in the latter 0.051, t=6.14, (Amphibia: Anura) on New Guinea: A mitochondrial phyloge- P=0.000007) (Fig. 12). There are continuous dorsolater- ny reveals parallel evolution of morphological and life histo- al skin ridges in C. tetzlaffi, but discontinuous dorsolat- ry traits and disproves the current morphology-based classi- fication. Molecular Phylogenetics and Evolution 47: 353–365 eral skin glands in C. monosyllabus sp. n. Kraus F, Allison A (2006) Three new species of Cophixalus (Anura: Microhylidae) from southeastern New Guinea. Her- Apart from these morphological differences, and most im- petologica 62: 202–220 portant for species differentiation, are the advertisement Kraus F, Allison A (2009) New species of Cophixalus (Anura: calls: C. monosyllabus sp. n. utters a single peeping note Microhylidae) from Papua New Guinea. Zootaxa 2128: 1–38 with a mean duration of 196 ms (range 173–224 ms), Misof B, Misof K. (2009) A Monte Carlo approach successful- while the call of C. tetzlaffi consists of three to four peep- ly identifies randomness in multiple sequence alignments: A more objective means of data exclusion. Systematic Biology ing notes with a mean note duration of 422 ms (range 58: 21–34 347–518 ms). Richards SJ, Oliver PM (2007) A new species of Cophixalus Acknowledgements. Field work and collection of voucher speci- (Anura: Microhylidae) from Misima Island, Papua New Gui- mens was permitted by representatives of Belai Besar Konser- nea. Pacific Science 61 (2): 279–287 vasi Sumber Daya Alam (KSDA), Sorong, Papua Province of Indonesia (PPI). Marthinus Kapisa (Biak/PPI), Andreas, Frank, and Apner Piahar (Kampung Lusiperi near Fakfak Town/PPI), and Christian Bergmann (Berlin/Germany) helped during field work. The Family Piahar also permitted the collection of frogs on their property. Rudolf Arndt (Pomona, New Jersey, USA) carefully read my draft and made a number of helpful comments. B. Stelbrink and T. von Rintelen (ZMB) “constructed” the mo- lecular tree. Elisa Forster (Zürich, Switzerland) prepared Figures Received: 27.VII.2010 2 and 3, and Frank Tillack (ZMB) provided technical help. To Accepted: 11.X.2010 all of them I am deeply grateful.

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